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Population Genetics of the Endemic Hawaiian Species Chrysodracon Hawaiiensis and Chrysodracon Auwahiensis (Asparagaceae): Insights from RAPD and ISSR Variation

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Population Genetics of the Endemic Hawaiian Species Chrysodracon Hawaiiensis and Chrysodracon Auwahiensis (Asparagaceae): Insights from RAPD and ISSR Variation International Journal of Molecular Sciences Article Population Genetics of the Endemic Hawaiian Species Chrysodracon hawaiiensis and Chrysodracon auwahiensis (Asparagaceae): Insights from RAPD and ISSR Variation Pei-Luen Lu 1,2,*, Mitsuko Yorkson 2 and Clifford W. Morden 2 1 Department of BioResources, Da-Yeh University, Changhua 51591, Taiwan 2 Department of Botany, University of Hawaii at Manoa,¯ Honolulu, HI 96822, USA; [email protected] (M.Y.); [email protected] (C.W.M.) * Correspondence: [email protected]; Tel.: +886-4-8511-888 (ext. 6215) Academic Editors: Jianhua Zhu and Marcello Iriti Received: 1 May 2016; Accepted: 11 August 2016; Published: 16 August 2016 Abstract: The genus Chrysodracon has six endemic species in the Hawaii Islands. Chrysodracon hawaiiensis is endemic to Hawaii Island and was described as a distinct species in 1980. It was listed as an endangered species on the International Union for the Conservation of Nature and Natural Resources (IUCN) Red List in 1997. This woody plant species was, at one time, common in exposed dry forests, but it became very rare due to grazing pressure and human development. The tree species Chrysodracon auwahiensis (C. auwahiensis), endemic to Maui and Molokai, still has large adult populations in dry lands of the islands, but unfortunately no regeneration from seed has been reported in those areas for many years. The two endemic species were examined using the molecular technique of random amplified polymorphic DNA (RAPD) and inter simple sequence repeats (ISSR) to determine the genetic structure of the populations and the amount of variation. Both species possess similar genetic structure. Larger and smaller populations of both species contain similar levels of genetic diversity as determined by the number of polymorphic loci, estimated heterozygosity, and Shannon’s index of genetic diversity. Although population diversity of Chrysodracon hawaiiensis (C. hawaiiensis) is thought to have remained near pre-disturbance levels, population size continues to decline as recruitment is either absent or does not keep pace with senescence of mature plants. Conservation recommendations for both species are suggested. Keywords: Chrysodracon; conservation; Hawaiian species; ISSR; Pleomele; population genetics; RAPD 1. Introduction The Hawaiian Islands include a high percentage of endemic species and are one of 25 biodiversity hotspots in the world [1–4]. Although many genera are species-rich in Hawaii and have special evolutionary histories, few of them have been studied in detail [4–6]. Many Pacific Island species, including those from the Hawaiian Islands, have a fragile existence. This is often due to their populations being scattered broadly within or across different islands and a limited genetic diversity due to their recent colonization, isolation from the source population, and/or the population size being restricted within island environments [4,5,7,8]. A consequence of this fragility has resulted in many endemic Hawaiian plant species having become endangered and the level of genetic diversity present becoming severely reduced compounding the problems for species recovery [9–11]. For example, the Hawaiian dry forests have been seriously reduced due to habitat loss from commercial or agricultural development and the spread of invasive plant and animal species [5,12]. Notably, more than 90% of Hawaiian dry forests are already lost [13] and 50% of the extant Hawaiian endemic flora is listed as Int. J. Mol. Sci. 2016, 17, 1341; doi:10.3390/ijms17081341 www.mdpi.com/journal/ijms Int. J. Mol. Sci. 2016, 17, 1341 2 of 16 endangered or rare in the International Union for the Conservation of Nature and Natural Resources (IUCN) [14] or by the US Fish and Wildlife Service (USFWS) [15]. Therefore, the study and conservation of genetic resources in populations, species, and ecosystems are essential to maintaining biodiversity and population dynamics. The Hawaiian endemic genus Chrysodracon (Jankaski) Lu and Morden (Asparagaceae), species previously included among the widely distributed tropical genus Pleomele Salisbury, has six endemic species in the Hawaiian Islands [16,17]. Chrysodracon hawaiiensis (Degener and Degener) Lu and Morden was distinguished as a species in 1980 [18]. Unfortunately, populations of this species have declined rapidly in the past few decades and the USFWS listed C. hawaiiensis (as Pleomele hawaiiensis) as an endangered species in 1996 [15]. The IUCN also placed it on their red list of endangered and threatened species in 1997 [14]. Chrysodracon hawaiiensis exists in only 6–8 populations totaling approximately 300–400 individuals in sunny dry forests on the leeward side of Hawaii Island [15] (Figure1). The largest extant wild population with approximately 200 individuals is located at Puu Waawaa [15]. This species has a unique ability to grow in young lava substrates often on steep slopes. To date, nothing is known about the genetic structure of this species. Due to its rarity and small population sizes, it may possibly go extinct or become more severely restricted in distribution within the next few decades if appropriate conservation management are not adopted. Presently, C. hawaiiensis is the only species of Chrysodracon recognized as occurring on Hawaii Island [17]. However, St. John [19] had previously recognized three distinct species (within Pleomele) based on morphological differences: P. hawaiiensis (sensu stricto), P. kaupulehuensis St. John, and P. konaensis St. John. These three species were distinguished by leaf width and the perianth tube length. The perianth of P. hawaiiensis is 37–40 mm long with a perianth tube longer than 26 mm whereas the perianth of P. konaensis is less than 37 mm, with a perianth tube less than 23 mm, and the perianth of P. kaupulehuensis is greater than 43 mm long. The leaf width of P. hawaiiensis and P. konaensis is less than 22 mm compared to the leaf width of P. kaupulehuensis being greater than 23 mm [19]. The most recent treatment of these species combined them within the single species P. hawaiiensis (sensu lato)[17]. As such, it is also important to examine their population differentiation and genetic variation to gain a better understanding of their interrelations. There were two objectives of this study. First, to investigate the genetic structure within and among populations of the endangered species C. hawaiiensis. In doing so, comparisons will also be made of the level of diversity within populations of different size. Understanding the population genetic structure of the endemic Hawaiian Chrysodracon species will be desirable to provide the insight needed for proper conservation strategies to preserve the biodiversity of island ecosystems, reveal the evolutionary stages of those species, and address genetic resource problems that those populations are facing. It will also provide appropriate recovery suggestions for collecting the seeds and artificial pollination from those populations to incorporate the maximum genetic variation in these efforts. To best examine the population structure of an endangered species, it is also necessary to analyze the population structure of a non-endangered congener species for comparison. Therefore, our second objective was to conduct a genetic survey of Chrysodracon auwahiensis (Lu & Morden), a non-endangered species endemic to Maui (Figure1) and Molokai Islands, to estimate the level and distribution of genetic diversity among populations. There are several extant populations of C. auwahiensis with thousands of individuals present. After completing the population genetics study of both C. hawaiiensis and C. auwahiensis, a comparison between them will provide an understanding of the type of variation that possibly was present in populations of C. hawaiiensis prior to habitat degradation and alteration. Knowledge of the population structure and level of variation will assist in formulating management practices for this species. Int. J. Mol. Sci. 2016, 17, 1341 3 of 16 Int. J. Mol. Sci. 2016, 17, 1341 3 of 16 Figure 1. Locations of populations used in Chrysodracon genetic studies. (A) The eight Hawaiian Figure 1. Locations of populations used in Chrysodracon genetic studies. (A) The eight Hawaiian Islands; (B) topographic map of Hawaii Island; (C) topographic map of Maui Island. Populations Islands; (B) topographic map of Hawaii Island; (C) topographic map of Maui Island. Populations are are demarcated with the color used in the principal coordinate analysis (PCO) analysis. Names of demarcated with the color used in the principal coordinate analysis (PCO) analysis. Names of volcanoes on each island are indicated for Hawaii (5) and Maui (2); principle cities associated with each volcanoes on each island are indicated for Hawaii (5) and Maui (2); principle cities associated with islands are indicated with a star ( ). each islands are indicated with a starF (). 2.2. Results Results InIn general,general, thethe genetic genetic diversity diversity measures measures in both in both random random amplified amplified polymorphic polymorphic DNA (RAPD) DNA (RAPD)and inter and simple inter sequence simple sequence repeat (ISSR) repeat analyses (ISSR) wereanalyses very were similar very and similar results and obtained results were obtained highly werecompatible. highly Overall,compatible. the geneticOverall, diversity the genetic values diversity were lower values in RAPDwere lower than ISSRin RAPD analyses than and ISSR the analysesvalues for and population the values differentiation
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